Long-term trends in cetacean occurrence during the annual sardine run off the Wild Coast,South Africa

During the austral winter, cetaceans and other apex predators follow the annual northeastward movement of shoaling sardines, known as the sardine run, along the southeast coast of South Africa, including a 400-km stretch called the Wild Coast. In total, 131 opportunistic aerial surveys were conducted between May and July, from 1996 to 2014, to monitor sardine movement. Cetacean-sighting data from these surveys were analysed, focusing on long-term trends in frequencies of the cetaceans. In total, 630 sightings involving five cetacean species were recorded: 268 (approximately 32 400 individuals) of Indo-Pacific bottlenose dolphin Tursiops aduncus, 108 (approximately 79 400 individuals) of long-beaked common dolphin Delphinus capensis, 242 (approximately 670 individuals) of humpback whale Megaptera novaeangliae, 1 (two individuals) of southern right whale Eubalaena australis, and 11 (16 individuals) of Bryde’s whale Balaenoptera edeni. The occurrence of common dolphins, typically associated with sardines, decreased significantly in average group size over the study period (p = 0.0343); bottlenose dolphins, considered generalist feeders, demonstrated no such trend (p = 0.916). Humpback whales were most frequently sighted between 2010 and 2014, and with significantly larger groups observed towards the end of the study period (p = 0.0121). For all these species, more sightings were made inside than outside of marine protected areas (>70% of the dolphin species, and >65% of the humpback whales), both pre- and post-2005 (from 2005 the size of the survey area increased). The results indicate that movements of the common dolphin may be employed as a proxy for sardine occurrence. Long-term trends evident in the data also demonstrate the importance of this coastal region for bottlenose dolphins as well as use as a migratory corridor for humpback whales.     

Sharks caught in the KwaZulu-Natal bather protection programme,South Africa. 14. The smooth hammerhead shark Sphyrna zygaena (Linnaeus)

The current study provides long-term catch-rate, biological and feeding data for smooth hammerhead sharks, Sphyrna zygaena, caught in South Africa’s KwaZulu-Natal bather protection programme. In total, 2 512 S. zygaena were caught in net installations between 1978 and 2014, and 72 S. zygaena were caught on drumlines between 2007 and 2014. There was no significant log-linear year trend in the net catch rate over time (slope = 0.0054, t = 1.808, p = 0.07). However, there was a significant temporal increase in mean size of the captured sharks (slope = 0.0012, t = 3.502, p < 0.001). A quasi-Poisson generalised additive mixed model showed that increasing latitude, winter months, colder sea temperatures and the deployment of drumlines all had a significant positive effect on the catch rate of sharks in nets. The size frequency of the catch was unimodal, with significantly more females caught in the nets and more males on the drumlines. The majority (93.1%) of all sharks caught were immature and measured between 80 and 120 cm precaudal length. Teleosts and cephalopods dominated the sharks’ diet in terms of all dietary indices. The prey species consumed indicate that immature S. zygaena are feeding primarily within the pelagic zone of shallow coastal habitats.     

The 2005 KwaZulu-Natal sardine run survey sheds new light on the ecology of small pelagic fish off the east coast of South Africa

Despite much public awareness surrounding the annual migration of sardine Sardinops sagax northward along the east coast of South Africa in winter each year, relatively little research effort has been expended to improve understanding of the ‘sardine run’. For this reason, a dedicated multidisciplinary survey, timed to coincide with the annual sardine run, was conducted off the East Coast in June and July of 2005. The major objective of the survey was to estimate the biomass of sardine off the East Coast during the run, and to compare this with biomass estimates collected during previous surveys conducted in this area during the late 1980s when the South African sardine population was at a considerably smaller size. We also collected data on the distribution of sardine and other small pelagic fish species and their eggs, the biological characteristics of sardine during the run, and data on the hydrography (temperature and currents) and lower trophic levels (phytoplankton and zooplankton) of the region. Results suggest that the biomass of sardine off the East Coast in winter remains relatively small and consistent, regardless of overall sardine population size. The narrow continental shelf to the east of Port Alfred, which is dominated offshore by the fast-flowing warm Agulhas Current, constrains the amount of suitable habitat for sardine and other clupeoids such as anchovy Engraulis encrasicolus, West Coast round herring Etrumeus whiteheadi and East Coast round herring Etrumeus teres, and hence precludes these species from attaining a high biomass in this region. Additionally, primary and secondary productivity levels are much lower than elsewhere on the western and eastern Agulhas Bank off the south coast of South Africa, suggesting that the sardine run is not a feeding migration. A previous hypothesis that the run is mainly a result of an expansion of the distributional range of these fish as conditions become favourable in winter due to sporadic cooling off the East Coast is also not entirely supported by results from the survey. It is suggested that a migration for the purposes of spawning off this coast when conditions become favourable is a more likely incentive for sardine to undertake this arduous journey, despite increased predation and poor feeding conditions.     

Sharks caught in the protective gill nets off KwaZulu-Natal,South Africa. 12. The African angel shark Squatina africana (Regan)

Between 1980 and 2001, a total of 661 African angel sharks Squatina africana was caught in the protective nets off KwaZulu-Natal, South Africa. The mean annual catch was 30 sharks (range = 11–69, SD = 12.4), with no trend in catch rate over the study period. Individuals were caught throughout the year and through much of the netted region, with a higher catch to the south. The sex ratio of the catch was 2.9 females:1 male. Males matured between 640mm and 700mm and females at about 700mm precaudal length (PCL). Mature males had clusters of thorn-shaped denticles, each about 2mm high, near the anterio-dorsal margins of both pectoral and pelvic fins. Of the mature females, 44% were pregnant, many of which contained only ova in utero. Embryos were present from April through to January. The average litter size was six, with length at parturition at least 240mm PCL. Most early-term pregnant females and all mature males were caught in the south. The reproductive cycle showed some seasonality and appeared to be biennial, with a gestation of about one year. Teleosts were the most common prey (76% of stomachs with food), followed by cephalopods (52%).     

Sharks caught in the protective gill nets off KwaZulu-Natal,South Africa. 11. The scalloped hammerhead shark Sphyrna lewini (Griffith and Smith)

Between 1978 and 1998, a total of 3 385 scalloped hammerhead sharks Sphyrna lewini was caught in the protective nets off KwaZulu-Natal. The mean annual catch was 166 sharks (range 60–279). There was a significant decrease in catch rate with time, but the relationship with the population size in KwaZulu-Natal waters is unknown. Size and sex segregation is indicated. Sizes-at-50% maturity were 161.5cm (males) and 183.1cm (females). Most of the catch (91%) was immature, but neonates are poorly sampled by the 51-cm meshed nets. In small animals (<160cm precaudal length, PCL), males significantly outnumbered females by 2.2:1, and in large animals by 3.6:1. The length-mass relationship differed between sexes. Catches of both small and large animals were highest in summer. Most of the large males were caught in November and December, consistent with an inshore movement of mature animals to breed in summer, but no evidence of recent mating was observed in either sex. Females pregnant with term embryos (median embryo length per litter 30.4–36.2cm) were caught between October and March. These females tended to have maximum ovarian follicle diameters of ≥30mm, indicating that mating would occur shortly after parturition. The median size of 11 litters was 10 embryos. The Tugela Bank to the north of the netted region appears to be a nursery ground. Teleosts, comprising 42 families and 60 identified species, dominated the diet in terms of frequency of occurrence (77%), followed by cephalopods (25%).     

Shelf currents,lee-trapped and transient eddies on the inshore boundary of the Agulhas Current,South Africa: their relevance to the KwaZulu-Natal sardine run

The existence and strength of the annual KwaZulu-Natal (KZN) sardine run has long been a conundrum to fishers and scientists alike ― particularly that the sardine Sardinops sagax migrate along the narrow Transkei shelf against the powerful, warm Agulhas Current. However, examination of ship-borne acoustic Doppler current profiler (S–ADCP) data collected during two research surveys in 2005 indicated that northward-flowing coastal countercurrents exist at times between the Agulhas Bank and the KZN Bight, near Port Alfred, East London, Port St Johns and Durban. The countercurrent near Port Alfred extended as far east as the Keiskamma River, within an upwelling zone known to exist there. An ADCP mooring at a depth of 32 m off Port Alfred indicated that the countercurrent typically lasted a few days, but at times remained in the same direction for as long as 10 days. Velocities ranged between 20 and 60 cm s^?1 with maximum values of ～80 cm s^?1. The S–ADCP data also highlighted the existence of cyclonic flow in the Port St Johns–Waterfall Bluff coastal inset, with a northward coastal current similarly ranging in velocity between 20 and 60 cm s^?1. CTD data indicated that this was associated with shelf-edge upwelling, with surface temperatures 2–4 °C cooler than the adjacent core temperature (24–26 °C) of the Agulhas Current. Vertical profiles of the S–ADCP data showed that the countercurrent, about 7 km wide, extends down the slope to at least 600 m, where it appeared to link with the deep Agulhas Undercurrent at 800 m. S–ADCP and sea surface temperature (SST) satellite data confirmed the existence of the semi-permanent, lee-trapped, cyclonic eddy off Durban, associated with a well-defined northward coastal current between Park Rynie and Balito Bay. Analysis of three months (May–July 2005) of satellite SST and ocean colour data showed the shoreward core-boundary of the Agulhas Current (24 °C isotherm) to commonly be close to the coast along the KZN south coast, as well as between the Kei and Mbhashe rivers on the Transkei shelf. The Port St Johns–Waterfall Bluff cyclonic eddy was also frequently visible in these satellite data. Transient cyclonic eddies, which spanned 150–200 km of shelf, appeared to move downstream in the shoreward boundary of the Agulhas Current at a frequency of about once a month. These seemed to be break-away Durban eddies. Data collected by ADCP moorings deployed off Port Edward in 2005 showed that these break-away eddies and the well-known Natal Pulse are associated with temporary northward countercurrents on the shelf, which can last up to six days. It is proposed that these countercurrents off Port Alfred, East London and Port St Johns assist sardine to swim northwards along the Transkei shelf against the Agulhas Current, but that their progress north of Waterfall Bluff is dependent on the arrival of a transient, southward-moving, break-away Durban cyclonic eddy, which apparently sheds every 4–6 weeks, or on the generation of a Natal Pulse. This passage control mechanism has been coined the ‘Waterfall Bluff gateway’ hypothesis. The sardine run survey in June–July 2005 was undertaken in the absence of a cyclonic eddy on the KZN south coast, i.e. when the ‘gate’ was closed.